To burn, or not to burn: that is the question.

Recently, Zylstra et al. reported that wet sclerophyll forest left unburnt for 75 years experiences a marked decrease in flammability, requiring a radical rethink about fire management. This also highlights the vertical dimension of fires, with species conservation favored by a mosaic of fire types (high pyrodiversity).

Fossil pollen resolves origin of the South African Proteaceae as transcontinental not transoceanic.

BACKGROUND AND AIMS: The prevailing view from the areocladogenesis of molecular phylogenies is that the iconic South African Cape Proteaceae (subfamily Proteoideae) arrived from Australia across the Indian Ocean in the Upper Cretaceous (100‒65 million years ago, Ma). Since fossil pollen indicates that the family probably arose in North-West Africa during the early Cretaceous, an alternative view is that it migrated to the Cape from North-Central Africa. The plan therefore was to collate fossil pollen records throughout Africa to determine if they are consistent with an African (para-autochthonous) origin for the Cape Proteaceae, and to seek further support from other paleo-disciplines. METHODS: Palynology (identity, date and location of records), molecular phylogeny and chronogram preparation, biogeography of plate tectonics, and paleo-atmospheric and ocean circulation models. KEY RESULTS: Our collation of the rich assemblage of Proteaceae palynomorphs stretching back to 107 Ma (Triorites africaensis) in North-West Africa showed its progressive overland migration to the Cape by 75‒65 Ma. No key palynomorphs recorded in Australia-Antarctica have morphological affinities with African fossils but specific clade assignment of the preMiocene records is not currently possible. The Cape Proteaceae encompass three molecular-based clades (tribes) whose most-recent apparent ancestors are sisters to those in Australia. However, our chronogram shows that the major Adenanthos/Leucadendron-related clade, originating 54‒34 Ma, would have 'arrived' too late as species with Proteaceae affinities were already present ~20 My earlier. The Franklandia/Protea-related clade arose 118‒81 Ma so its distinctive pollen should have been the foundation for the scores of palynomorphs recorded at 100‒80 Ma but it was not. Also, the prevailing winds and ocean currents trended away from South Africa rather than towards, as the 'out-of-Australia' hypothesis requires. Based on the evidence assembled here, we list three points favouring an Australian origin and nine against; four points favouring an Antarctic origin and seven against; and nine points favouring a North-Central African origin and three against. CONCLUSIONS: We conclude that a gradual migration of the Proteaceae from North-Central Africa southeast→south→southwest to the Cape and surrounds occurred via adaptation and speciation during the period 90‒70 Ma. We caution that incorrect conclusions may be drawn from literal interpretations of molecular phylogenies that neglect the fossil record and do not recognize the possible confounding effects of selection under matched environments leading to parallel evolution and extinction of bona fide sister clades.

Relative growth rate (RGR) and other confounded variables: mathematical problems and biological solutions.

BACKGROUND: Relative growth rate (RGR) has a long history of use in biology. In its logged form, RGR = ln[(M + ΔM)/M], where M is size of the organism at the commencement of the study, and ΔM is new growth over time interval Δt. It illustrates the general problem of comparing non-independent (confounded) variables, e.g. (X + Y) vs. X. Thus, RGR depends on what starting M(X) is used even within the same growth phase. Equally, RGR lacks independence from its derived components, net assimilation rate (NAR) and leaf mass ratio (LMR), as RGR = NAR × LMR, so that they cannot legitimately be compared by standard regression or correlation analysis. FINDINGS: The mathematical properties of RGR exemplify the general problem of 'spurious' correlations that compare expressions derived from various combinations of the same component terms X and Y. This is particularly acute when X >> Y, the variance of X or Y is large, or there is little range overlap of X and Y values among datasets being compared. Relationships (direction, curvilinearity) between such confounded variables are essentially predetermined and so should not be reported as if they are a finding of the study. Standardizing by M rather than time does not solve the problem. We propose the inherent growth rate (IGR), lnΔM/lnM, as a simple, robust alternative to RGR that is independent of M within the same growth phase. CONCLUSIONS: Although the preferred alternative is to avoid the practice altogether, we discuss cases where comparing expressions with components in common may still have utility. These may provide insights if (1) the regression slope between pairs yields a new variable of biological interest, (2) the statistical significance of the relationship remains supported using suitable methods, such as our specially devised randomization test, or (3) multiple datasets are compared and found to be statistically different. Distinguishing true biological relationships from spurious ones, which arise from comparing non-independent expressions, is essential when dealing with derived variables associated with plant growth analyses.

Extrafloral nectar as entrée and elaiosomes as main course for ant visitors to a fireprone, mediterranean-climate shrub.

Thousands of plants produce both extrafloral nectaries (EFNs) on their leaves and nutrient-rich appendages on their diaspores (elaiosomes). Although their individual ecology is well-known, any possible functional link between these structures has almost always been ignored. Here, we recognized their co-presence in the shrub, Adenanthos cygnorum (Proteaceae), and studied their function and interaction. We observed that the same ants frequently visit both structures, seeds are attractive to vertebrate granivores but are released into a leafy cup from where they are harvested by ants and taken to their nests, from which seeds, lacking elaiosomes, germinate after fire. We showed that juvenile plants do not produce EFNs and are not visited by ants. We conclude that EFNs are not just an indirect adaptation to minimize herbivory via aggressive ant visitors (the role of a minority) but specifically enhance reproductive success in two ways: First, by inducing ants to visit the plant as a reliable food source throughout the year. Second, by promoting discovery of the seasonally available, elaiosome-bearing seeds for transport to their nests (the majority of visitors), so avoiding the risk of granivory should seeds instead fall to the ground. Parasitoid wasps play a supporting role in controlling the main insect herbivore whose larvae devour the reproductive apices. Thus, the EFN-elaiosome relationship has three components that enhance species fitness: foliage protection, seed transport, and granivore escape. A similar system has been described only once before (in an unrelated biome) and, consistent with the objectives of ecology as an integrative science, deserves wider study.

Fossil flowers of Phylica support a 250 Ma origin for Rhamnaceae.

A new fossil discovery reported by Shi et al. changes our understanding of the biogeographic history of the cosmopolitan family, Rhamnaceae. Flowering shoots of the African genus Phylica (Rhamnaceae) dated at 100 million years ago (Ma) imply a 250 Ma origin of the family in fire-prone Gondwanan vegetation that enabled overland dispersal to all continents where it is currently widespread.

Ancient Rhamnaceae flowers impute an origin for flowering plants exceeding 250-million-years ago.

Setting the molecular clock to newly described 100-million-year-old flowering shoots of Phylica in Burmese amber enabled us to recalibrate the phylogenetic history of Rhamnaceae. We traced its origin to ∼260 million years ago (Ma) that can explain its migration within and beyond Gondwana since that time and implies an origin for flowering plants that stretches well beyond 290 Ma. Ancestral trait assignments also revealed that hard-seededness, fire-proneness, and to a lesser extent, heat-released seed dormancy, have a similarly long history in this clade.

Fire-released seed dormancy - a global synthesis.

Seed dormancy varies greatly between species, clades, communities, and regions. We propose that fireprone ecosystems create ideal conditions for the selection of seed dormancy as fire provides a mechanism for dormancy release and postfire conditions are optimal for germination. Thus, fire-released seed dormancy should vary in type and abundance under different fire regimes. To test these predictions, we compiled data from a wide range of fire-related germination experiments for species in different ecosystems across the globe. We identified four dormancy syndromes: heat-released (physical) dormancy, smoke-released (physiological) dormancy, non-fire-released dormancy, and non-dormancy. In fireprone ecosystems, fire, in the form of heat and/or chemical by-products (collectively termed 'smoke'), are the predominant stimuli for dormancy release and subsequent germination, with climate (cold or warm stratification) and light sometimes playing important secondary roles. Fire (heat or smoke)-released dormancy is best expressed where woody vegetation is dense and fires are intense, i.e. in crown-fire ecosystems. In such environments, seed dormancy allows shade-intolerant species to take advantage of vegetation gaps created by fire and synchronize germination with optimal recruitment conditions. In grassy fireprone ecosystems (e.g. savannas), where fires are less intense but more frequent, seed dormancy is less common and dormancy release is often not directly related to fire (non-fire-released dormancy). Rates of germination, whether controls or postfire, are twice as fast in savannas than in mediterranean ecosystems. Fire-released dormancy is rare to absent in arid ecosystems and rainforests. The seeds of many species with fire-released dormancy also possess elaiosomes that promote ant dispersal. Burial by ants increases insulation of seeds from fires and places them in a suitable location for fire-released dormancy. The distribution of these dormancy syndromes across seed plants is not random - certain dormancy types are associated with particular lineages (phylogenetic conservatism). Heat-released dormancy can be traced back to fireprone floras in the 'fiery' mid-Cretaceous, followed by smoke-released dormancy, with loss of fire-related dormancy among recent events associated with the advent of open savannas and non-fireprone habitats. Anthropogenic influences are now modifying dormancy-release mechanisms, usually decreasing the role of fire as exaptive effects. We conclude that contrasting fire regimes are a key driver of the evolution and maintenance of diverse seed dormancy types in many of the world's natural ecosystems.

Fire-mediated germination syndromes in Leucadendron (Proteaceae) and their functional correlates.

A mechanistic understanding of fire-driven seedling recruitment is essential for effective conservation management of fire-prone vegetation, such as South African fynbos, especially with rare and threatened taxa. The genus Leucadendron (Proteaceae) is an ideal candidate for comparative germination studies, comprising 85 species with a mixture of contrasting life-history traits (killed by fire vs able to resprout; serotinous vs geosporous) and seed morphologies (nutlets vs winged achenes). Individual and combined effects of heat and smoke on seed germination of 40 species were quantified in the laboratory, and Bayesian inference applied to distinguish biologically meaningful treatment effects from non-zero, but biologically trivial, effects. Three germination syndromes were identified based on whether germination was dependent on, enhanced by, or independent of direct fire cues (heat and smoke). Seed storage location was the most reliable predictor of germination syndromes, with soil-stored seeds c. 80% more likely to respond to direct fire cues (primarily smoke) than canopy-stored seeds. Notable exceptions were L. linifolium, with an absolute requirement for smoke to germinate (the third serotinous species so reported), and two other serotinous species with smoke-enhanced germination. Nutlet-bearing species, whether serotinous or geosporous, were c. 70% more likely to respond to fire cues than winged seeds, but there was no evidence for an effect of phylogeny or persistence strategy on germination. This comprehensive account of seed germination characteristics and identification of germination syndromes and their predictors, supports propagation, conservation and restoration initiatives in this iconic fynbos genus and other fire-prone shrubs with canopy or soil-stored seeds.

Plant functional types determine how close postfire seedlings are from their parents in a species-rich shrubland.

BACKGROUND AND AIMS: Fine-scale spatial patterns of the seedlings of co-occurring species reveal the relative success of reproduction and dispersal and may help interpret coexistence patterns of adult plants. To understand whether postfire community dynamics are controlled by mathematical, biological or environmental factors, we documented seedling-adult (putative parent) distances for a range of co-occurring species. We hypothesized that nearest-seedling-to-adult distances should be a function of the distance between the closest conspecific seedlings, closest inter-adult distances and seedling-to-parent ratios, and also that these should scale up in a consistent way from all individuals, to within and between species and finally between functional types (FTs). METHODS: We assessed seedling-adult, seedling-seedling and adult-adult distances for 19 co-occurring shrub species 10 months after fire in a species-rich shrubland in south-western Australia. Species were categorized into 2 × 2 FTs: those that are killed by fire [non-(re)sprouters] vs. those that survive (resprouters) in nine taxonomically matched pairs, and those that disperse their seeds prefire (geosporous) vs. those that disperse their seeds postfire (serotinous). KEY RESULTS: For the total data set and means for all species, seedling-adult distance was essentially a mathematical phenomenon, and correlated positively with seedling-seedling distance and adult-adult distance, and inversely with seedlings per adult. Among the four FTs, seedling-adult distance was shortest for geosporous non-sprouters and widest for serotinous resprouters. Why adults that produce few seedlings (resprouters) should be further away from them defies a simple mathematical or biological explanation at present. Ecologically, however, it is adaptive: the closest seedling was usually under the (now incinerated) parent crown of non-sprouters whereas those of resprouters were on average four times further away. CONCLUSIONS: Our study highlights the value of recognizing four reproductive syndromes within fire-prone vegetation, and shows how these are characterized by marked differences in their seedling-adult spatial relations that serve to enhance biodiversity of the community.

Fire as a key driver of Earth's biodiversity.

Many terrestrial ecosystems are fire prone, such that their composition and structure are largely due to their fire regime. Regions subject to regular fire have exceptionally high levels of species richness and endemism, and fire has been proposed as a major driver of their diversity, within the context of climate, resource availability and environmental heterogeneity. However, current fire-management practices rarely take into account the ecological and evolutionary roles of fire in maintaining biodiversity. Here, we focus on the mechanisms that enable fire to act as a major ecological and evolutionary force that promotes and maintains biodiversity over numerous spatiotemporal scales. From an ecological perspective, the vegetation, topography and local weather conditions during a fire generate a landscape with spatial and temporal variation in fire-related patches (pyrodiversity), and these produce the biotic and environmental heterogeneity that drives biodiversity across local and regional scales. There have been few empirical tests of the proposition that 'pyrodiversity begets biodiversity' but we show that biodiversity should peak at moderately high levels of pyrodiversity. Overall species richness is greatest immediately after fire and declines monotonically over time, with postfire successional pathways dictated by animal habitat preferences and varying lifespans among resident plants. Theory and data support the 'intermediate disturbance hypothesis' when mean patch species diversity is correlated with mean fire intervals. Postfire persistence, recruitment and immigration allow species with different life histories to coexist. From an evolutionary perspective, fire drives population turnover and diversification by promoting a wide range of adaptive responses to particular fire regimes. Among 39 comparisons, the number of species in 26 fire-prone lineages is much higher than that in their non-fire-prone sister lineages. Fire and its byproducts may have direct mutagenic effects, producing novel genotypes that can lead to trait innovation and even speciation. A paradigm shift aimed at restoring biodiversity-maintaining fire regimes across broad landscapes is required among the fire research and management communities. This will require ecologists and other professionals to spread the burgeoning fire-science knowledge beyond scientific publications to the broader public, politicians and media.

Environmental drivers and genomic architecture of trait differentiation in fire-adapted Banksia attenuata ecotypes.

Trait divergence between populations is considered an adaptive response to different environments, but to what extent this response is accompanied by genetic differentiation is less clear since it may be phenotypic plasticity. In this study, we analyzed phenotypic variation between two Banksia attenuata growth forms, lignotuberous (shrub) and epicormic resprouting (tree), in fire-prone environments to identify the environmental factors that have driven this phenotypic divergence. We linked genotype with phenotype and traced candidate genes using differential gene expression analysis. Fire intervals determined the phenotypic divergence between growth forms in B. attenuata. A genome-wide association study identified 69 single nucleotide polymorphisms, putatively associated with growth form, whereas no growth form- or phenotype-specific genotypes were identified. Genomic differentiation between the two growth forms was low (Fst = 0.024). Differential gene expression analysis identified 37 genes/transcripts that were differentially expressed in the two growth forms. A small heat-shock protein gene, associated with lignotuber presence, was differentially expressed in the two forms. We conclude that different fire regimes induce phenotypic polymorphism in B. attenuata, whereas phenotypic trait divergence involves the differential expression of a small fraction of genes that interact strongly with the disturbance regime. Thus, phenotypic plasticity among resprouters is the general strategy for surviving varying fire regimes.

Evolutionary history of fire-stimulated resprouting, flowering, seed release and germination.

Fire has shaped the evolution of many plant traits in fire-prone environments: fire-resistant tissues with heat-insulated meristems, post-fire resprouting or fire-killed but regenerating from stored seeds, fire-stimulated flowering, release of on-plant-stored seeds, and germination of soil-stored seeds. Flowering, seed release and germination fit into three categories of response to intensifying fire: fire not required, weakly fire-adapted or strongly fire-adapted. Resprouting also has three categories but survival is always reduced by increasing fire intensity. We collated 286 records for 20 angiosperm and two gymnosperm families and 50 trait assignments to dated phylogenies. We placed these into three fire-adapted trait types: those associated with the origin of their clade and the onset of fire-proneness [primary diversification, contributing 20% of speciation events over the last 120 million years (My)], those originating much later coincident with a change in the fire regime (secondary diversification, 30%), and those conserved in the daughter lineage as already adapted to the fire regime (stabilisation, 50%). All four fire-response types could be traced to >100 My ago (Mya) with pyrogenic flowering slightly younger because of its dependence on resprouting. There was no evidence that resprouting was always an older trait than either seed storage or non-sprouting throughout this period, with either/both ancestral or derived in different clades and times. Fire-adapted traits evolved slowly in the Cretaceous, 120-65 Mya, and rapidly but fitfully in the Cenozoic, 65-0 Mya, peaking over the last 20 My. The four trait-types climaxed at different times, with the peak in resprouter speciation over the last 5 My attributable to fluctuating growing conditions and increasing savanna grasslands unsuitable for non-sprouters. All experienced a trough in the 40-30-Mya period following a reduction in world temperatures and oxygen levels and expected reduced fire activity. Thick bark and serotiny arose in the Mid-Cretaceous among extant Pinaceae. Heat-stimulated germination of hard seeds is ancestral in the 103-My-old Fabales. Smoke-(karrikin)-stimulated germination of non-hard seeds is even older, and includes the 101-My-old Restionaceae-Anarthriaceae. A smoke/karrikin response is detectable in some fire-free lineages that prove to have a fire-prone ancestry. Among clades that are predominantly fire-prone, absence of fire-related traits is the advanced condition, associated either with increased fire frequency (loss of serotiny and soil storage), or migration to fire-free habitats (loss of thick bark, pyrogenic flowering, serotiny or soil storage). Protea (Africa) and Hakea (Australia) illustrate the importance of stabilisation processes between resprouting/non-sprouting in accounting for speciation events over the last 20 My and highlight the frequent interchange possible between these two traits. Apart from Pinus, most ancestral trait reconstruction relative to fire has been conducted on predominantly Southern Hemisphere clades and this needs to be redressed. Despite these limitations, it is clear that fire has had a profound effect on fire-related trait evolution worldwide, and set the platform for subsequent evolution of many non-fire-related traits. Genetics of the triggering mechanisms remain poorly understood, except the karrikin system for smoke-stimulated germination. We exhort biologists to include fire-proneness and fire-related traits in their thinking on possible factors controlling the evolution of plants.

Fire and Plant Diversification in Mediterranean-Climate Regions.

Despite decades of broad interest in global patterns of biodiversity, little attention has been given to understanding the remarkable levels of plant diversity present in the world's five Mediterranean-type climate (MTC) regions, all of which are considered to be biodiversity hotspots. Comprising the Mediterranean Basin, California, central Chile, the Cape Region of South Africa, and southwestern Australia, these regions share the unusual climatic regime of mild wet winters and warm dry summers. Despite their small extent, covering only about 2.2% of world land area, these regions are home to approximately one-sixth of the world vascular plant flora. The onset of MTCs in the middle Miocene brought summer drought, a novel climatic condition, but also a regime of recurrent fire. Fire has been a significant agent of selection in assembling the modern floras of four of the five MTC regions, with central Chile an exception following the uplift of the Andes in the middle Miocene. Selection for persistence in a fire-prone environment as a key causal factor for species diversification in MTC regions has been under-appreciated or ignored. Mechanisms for fire-driven speciation are diverse and may include both directional (novel traits) and stabilizing selection (retained traits) for appropriate morphological and life-history traits. Both museum and nursery hypotheses have important relevance in explaining the extant species richness of the MTC floras, with fire as a strong stimulant for diversification in a manner distinct from other temperate floras. Spatial and temporal niche separation across topographic, climatic and edaphic gradients has occurred in all five regions. The Mediterranean Basin, California, and central Chile are seen as nurseries for strong but not spectacular rates of Neogene diversification, while the older landscapes of southwestern Australia and the Cape Region show significant components of both Paleogene and younger Neogene speciation in their diversity. Low rates of extinction suggesting a long association with fire more than high rates of speciation have been key to the extant levels of species richness.

Unearthing belowground bud banks in fire-prone ecosystems.

Despite long-time awareness of the importance of the location of buds in plant biology, research on belowground bud banks has been scant. Terms such as lignotuber, xylopodium and sobole, all referring to belowground bud-bearing structures, are used inconsistently in the literature. Because soil efficiently insulates meristems from the heat of fire, concealing buds below ground provides fitness benefits in fire-prone ecosystems. Thus, in these ecosystems, there is a remarkable diversity of bud-bearing structures. There are at least six locations where belowground buds are stored: roots, root crown, rhizomes, woody burls, fleshy swellings and belowground caudexes. These support many morphologically distinct organs. Given their history and function, these organs may be divided into three groups: those that originated in the early history of plants and that currently are widespread (bud-bearing roots and root crowns); those that also originated early and have spread mainly among ferns and monocots (nonwoody rhizomes and a wide range of fleshy underground swellings); and those that originated later in history and are strictly tied to fire-prone ecosystems (woody rhizomes, lignotubers and xylopodia). Recognizing the diversity of belowground bud banks is the starting point for understanding the many evolutionary pathways available for responding to severe recurrent disturbances.

Small-seeded Hakea species tolerate cotyledon loss better than large-seeded congeners.

Six Hakea species varying greatly in seed size were selected for cotyledon damage experiments. The growth of seedlings with cotyledons partially or completely removed was monitored over 90 days. All seedlings perished by the fifth week when both cotyledons were removed irrespective of seed size. Partial removal of cotyledons caused a significant delay in the emergence of the first leaf, and reduction in root and shoot growth of the large-seeded species. The growth of seedlings of small-seeded species was less impacted by cotyledon damage. The rate of survival, root and shoot lengths and dry biomass of the seedlings were determined after 90 days. When seedlings were treated with balanced nutrient solutions following removal of the cotyledons, survival was 95-98%, but 0% when supplied with nutrient solutions lacking N or P or with water only. The addition of a balanced nutrient solution failed to restore complete growth of any species, but the rate of root elongation for the small-seeded species was maintained. Cotyledons provide nutrients to support early growth of Hakea seedlings, but other physiological roles for the cotyledons are also implicated. In conclusion, small-seeded Hakea species can tolerate cotyledons loss better than large-seeded species.

Fire-Proneness as a Prerequisite for the Evolution of Fire-Adapted Traits.

Fire as a major evolutionary force has been disputed because it is considered to lack supporting evidence. If a trait has evolved in response to selection by fire then the environment of the plant must have been fire-prone before the appearance of that trait. Using outcomes of trait assignments applied to molecular phylogenies for fire-stimulated flowering, seed-release, and germination, in this Opinion article we show that fire-proneness precedes, or rarely coincides with, the evolution of these fire-adapted traits. In addition, fire remains central to understanding germination promoted by smoke among species occurring in non-fire-prone environments because of the historical association of their clade with fire. Fire-mimicking selection and associated exaptations have no place in understanding the evolution of fire-adapted traits because we find no support for any reversal in the fire-trait sequence through time.

A Cretaceous origin for fire adaptations in the Cape flora.

Fire has had a profound effect on the evolution of worldwide biotas. The Cape Floristic Region is one of the world's most species-rich regions, yet it is highly prone to recurrent fires and fire-adapted species contribute strongly to the overall flora. It is hypothesized that the current fire regimes in the Cape could be as old as 6-8 million years (My), while indirect evidence indicates that the onset of fire could have reached 18 million years ago (Ma). Here, we trace the origin of fire-dependent traits in two monocot families that are significant elements in the fire-prone Cape flora. Our analysis shows that fire-stimulated flowering originated in the Cape Haemodoraceae 81 Ma, while fire-stimulated germination arose in the African Restionaceae at least 70 Ma, implying that wildfires have been a significant force in the evolution of the Cape flora at least 60 My earlier than previous estimates. Our results provide strong evidence for the presence of fire adaptations in the Cape from the Cretaceous, leading to the extraordinary persistence of a fire-adapted flora in this biodiversity hotspot, and giving support to the hypothesis that Cretaceous fire was a global phenomenon that shaped the evolution of terrestrial floras.

Pre-Gondwanan-breakup origin of Beauprea (Proteaceae) explains its historical presence in New Caledonia and New Zealand.

New Caledonia and New Zealand belong to the now largely submerged continent Zealandia. Their high levels of endemism and species richness are usually considered the result of transoceanic dispersal events followed by diversification after they re-emerged from the Pacific Ocean in the mid-Cenozoic. We explore the origin and evolutionary history of Beauprea (Proteaceae), which is now endemic to New Caledonia but was once spread throughout eastern Gondwana, including New Zealand. We review the extensive Beauprea-type pollen data in the fossil records and analyze the relationship of these fossil taxa to extant genera within Proteaceae. We further reconstruct the phylogenetic relations among nine extant species of Beauprea and estimate the age of the Beauprea clade. By incorporating extinct taxa into the Beauprea phylogenetic tree, we reconstruct the ancient distribution of this genus. Our analysis shows that Beauprea originated c. 88 Ma (million years ago) in Antarctica-Southeastern Australia and spread throughout Gondwana before its complete breakup. We propose that Beauprea, already existing as two lineages, was carried with Zealandia when it separated from the rest of Gondwana c. 82 Ma, thus supporting an autochthonous origin for Beauprea species now in New Caledonia and historically in New Zealand up to 1 Ma. We show that the presence of Beauprea through transoceanic dispersal is implausible. This means that neither New Caledonia nor New Zealand has been entirely submerged since the Upper Cretaceous; thus, possible vicariance and allopatry must be taken into account when considering the high levels of endemism and species richness of these island groups.